|Male African leopard in Maasai Mara National Reserve, Kenya|
|Present and historical distribution of the leopard|
The leopard (Panthera pardus) is one of the five extant species in the genus Panthera, a member of the cat family, Felidae. It occurs in a wide range in sub-Saharan Africa, in some parts of Western and Central Asia, Southern Russia, and on the Indian subcontinent to Southeast and East Asia. It is listed as Vulnerable on the IUCN Red List because leopard populations are threatened by habitat loss and fragmentation, and are declining in large parts of the global range. The leopard is considered locally extinct in Hong Kong, Singapore, South Korea, Jordan, Morocco, Togo, the United Arab Emirates, Uzbekistan, Lebanon, Mauritania, Kuwait, Syria, Libya, Tunisia and most likely in North Korea, Gambia, Laos, Lesotho, Tajikistan, Vietnam and Israel. Contemporary records suggest that the leopard occurs in only 25% of its historical global range.
Compared to other wild cats, the leopard has relatively short legs and a long body with a large skull. Its fur is marked with rosettes. It is similar in appearance to the jaguar (Panthera onca), but has a smaller, lighter physique, and its rosettes are generally smaller, more densely packed and without central spots. Both leopards and jaguars that are melanistic are known as black panthers. The leopard is distinguished by its well-camouflaged fur, opportunistic hunting behaviour, broad diet, strength, and its ability to adapt to a variety of habitats ranging from rainforest to steppe, including arid and montane areas. It can run at speeds of up to 58 km/h (36 mph). The earliest known leopard fossils excavated in Europe are estimated 600,000 years old, dating to the late Early Pleistocene. Leopard fossils were also found in Sumatra, Taiwan and Japan.
The English name 'leopard' comes from Old French: leupart or Middle French: liepart, that derives from Latin: leopardus and Ancient Greek: λέοπάρδος (leopardos). Leopardos could be a compound of λέων (leōn), meaning lion, and πάρδος (pardos), meaning spotted. The word λέοπάρδος originally referred to a cheetah (Acinonyx jubatus).
'Panther' is another common name, derived from Latin: panther and Ancient Greek: πάνθηρ (pánthēr); The generic name Panthera originates in Latin: panthera, which refers to a hunting net for catching wild beasts that were used by the Romans in combats. Pardus is the masculine singular form.
The leopard's fur is generally soft and thick, notably softer on the belly than on the back. Its skin colour varies between individuals from pale yellowish to dark golden with dark spots grouped in rosettes. Its belly is whitish and its ringed tail is shorter than its body. Its pupils are round. Leopards living in arid regions are pale cream, yellowish to ochraceous and rufous in colour; those living in forests and mountains are much darker and deep golden. Spots fade toward the white underbelly and the insides and lower parts of the legs. Rosettes are circular in East African leopard populations, and tend to be squarish in Southern African and larger in Asian leopard populations. The fur tends to be grayish in colder climates, and dark golden in rain forest habitats. The pattern of the rosettes is unique in each individual. This pattern is thought to be an adaptation to dense vegetation with patchy shadows, where it serves as camouflage.
Its white-tipped tail is about 60–100 cm (23.6–39.4 in) long, white underneath and with spots that form incomplete bands toward the tail's end. The guard hairs protecting the basal hairs are short, 3–4 mm (0.1–0.2 in) in face and head, and increase in length toward the flanks and the belly to about 25–30 mm (1.0–1.2 in). Juveniles have woolly fur, and appear to be dark-coloured due to the densely arranged spots. Its fur tends to grow longer in colder climates. The leopard's rosettes differ from those of the jaguar (Panthera onca), which are darker and with smaller spots inside.
Size and weight
The leopard is sexually dimorphic with males larger and heavier than females. It is slender and muscular, with relatively short limbs and a broad head. Males stand 60–70 cm (23.6–27.6 in) at the shoulder, while females are 57–64 cm (22.4–25.2 in) tall. The head-and-body length ranges between 90 and 196 cm (2 ft 11.4 in and 6 ft 5.2 in) with a 66 to 102 cm (2 ft 2.0 in to 3 ft 4.2 in) long tail. Sizes vary geographically. Males weigh 37–90 kg (81.6–198.4 lb), and females 28–60 kg (61.7–132.3 lb). Some leopards in North Africa allegedly were as large as Barbary lions (Panthera leo leo). In 1913, an Algerian newspaper reported a leopard killed that allegedly measured about 275 cm (9 ft 0 in) in total length. The maximum weight of a wild leopard in Southern Africa was about 96 kg (212 lb). It measured 262 cm (8 ft 7.1 in). An Indian leopard killed in Himachal Pradesh in 2016 measured 261 cm (8 ft 6.8 in) with an estimated weight of 78.5 kg (173.1 lb); it was perhaps the largest known wild leopard in India.
The largest skull of a leopard was recorded in India in 1920 and measured 28 cm (11.0 in) in basal length, 20 cm (7.9 in) in breadth, and weighed 1,000 g (2 lb 4 oz). The skull of an African leopard measured 285.8 mm (11.25 in) in basal length, and 181.0 mm (7.125 in) in breadth, and weighed 790 g (1 lb 12 oz).
Melanistic leopards are also called black panthers. Melanism in leopards is caused by a recessive allele and inherited as a recessive trait. Interbreeding in melanistic leopards produces a significantly smaller litter size than is produced by normal pairings. The black leopard is common foremost in tropical and subtropical moist forests like the equatorial rainforest of the Malay Peninsula and the tropical rainforest on the slopes of some African mountains such as Mount Kenya. Between January 1996 and March 2009, leopards were photographed at 16 sites in the Malay Peninsula in a sampling effort of more than 1,000 camera trap nights. Of the 445 photographs of melanistic leopards, 410 were taken in study sites south of the Kra Isthmus, where the non-melanistic morph was never photographed. These data indicate the near-fixation of the dark allele in the region. The expected time for the fixation of this recessive allele due to genetic drift alone ranged from about 1,100 years to about 100,000 years. Pseudomelanistic leopards have also been reported.
In India, nine pale and white leopards were reported between 1905 and 1967. Leopards exhibiting erythrism were recorded between 1990 and 2015 in South Africa's Madikwe Game Reserve and in Mpumalanga. The cause of this morph known as a "strawberry leopard" or "pink panther" is not well understood.
Felis pardus was the scientific name proposed by Carl Linnaeus in 1758. The generic name Panthera was first used by Lorenz Oken in 1816, who included all the known spotted cats into this group. Oken's classification was not widely accepted, and Felis or Leopardus was used as the generic name until the early 20th century.
The leopard was designated as the type species of Panthera by Joel Asaph Allen in 1902. In 1917, Reginald Innes Pocock also subordinated the tiger (P. tigris), lion (P. leo), and jaguar (P. onca) to Panthera.
Following Linnaeus' first description, 27 leopard subspecies were proposed by naturalists between 1794 and 1956. Since 1996, only eight subspecies have been considered valid on the basis of mitochondrial analysis. Later analysis revealed a ninth valid subspecies, the Arabian leopard.
In 2017, the Cat Classification Task Force of the Cat Specialist Group recognized the following eight subspecies as valid taxa:
|African leopard (P. p. pardus) (Linnaeus, 1758)||It is the most widespread leopard subspecies and is native to most of Sub-Saharan Africa.|
|Indian leopard (P. p. fusca) (Meyer, 1794)||It is native to the Indian subcontinent, Myanmar and southern Tibet.|
|Javan leopard (P. p. melas) (Cuvier, 1809)||It is native to Java in Indonesia and is considered Critically Endangered.|
|Arabian leopard (P. p. nimr) (Hemprich and Ehrenberg, 1830)||It is native to the Arabian Peninsula, but considered locally extinct in the Sinai Peninsula. It is the smallest leopard subspecies.|
|P. p. tulliana (Valenciennes, 1856)||It is native to eastern Turkey, the Caucasus, southern Russia, the Iranian Plateau and the Hindu Kush. It is considered Endangered.|
|Amur leopard (P. p. orientalis) (Schlegel, 1857)||It is native to the Russian Far East and northern China, but is locally extinct in the Korean peninsula.|
|Indochinese leopard (P. p. delacouri) Pocock, 1930||It is native to mainland Southeast Asia and southern China.|
|Sri Lankan leopard (P. p. kotiya) Deraniyagala, 1956||It is native to Sri Lanka.|
Results of an analysis of molecular variance and pairwise fixation index of 182 African leopard museum specimens showed that some African leopards exhibit higher genetic differences than Asian leopard subspecies.
Results of phylogenetic studies based on nDNA and mtDNA analysis showed that the last common ancestor of the Panthera and Neofelis genera is thought to have lived about . Neofelis diverged about from the Panthera lineage. The tiger diverged about , followed by the snow leopard about and the leopard about . The leopard is a sister taxon to a clade within Panthera, consisting of the lion and the jaguar.
Results of a phylogenetic analysis of chemical secretions amongst cats indicated that the leopard is closely related to the lion. The geographic origin of the Panthera is most likely northern Central Asia. The leopard-lion clade was distributed in the Asian and African Palearctic since at least the early Pliocene. The leopard-lion clade diverged 3.1–1.95 million years ago. Additionally, a 2016 study revealed that the mitochondrial genomes of the leopard, lion and snow leopard are more similar to each other than their nuclear genomes, indicating that their ancestors hybridized with the snow leopard at some point in their evolution.
Fossils of leopard ancestors were excavated in East Africa and South Asia, dating back to the Pleistocene between 2 and . The modern leopard is suggested to have evolved in Africa about and to have radiated across Asia about 0.2 and . Fossil cat teeth collected in Sumatra's Padang Highlands were assigned to the leopard. It has since been hypothesized that it became extirpated on the island due to the Toba eruption about 75,000 years ago, and due to competition with the Sunda clouded leopard (Neofelis diardi) and the dhole (Cuon alpinus).
In Europe, the leopard occurred at least since the Pleistocene. Leopard-like fossil bones and teeth possibly dating to the Pliocene were excavated in Perrier in France, northeast of London, and in Valdarno, Italy. Until 1940, similar fossils dating back to the Pleistocene were excavated mostly in loess and caves at 40 sites in Europe, including Furninha Cave near Lisbon, Genista Caves in Gibraltar, and Santander Province in northern Spain to several sites across France, Switzerland, Italy, Austria, Germany, in the north up to Derby in England, in the east to Přerov in the Czech Republic and the Baranya in southern Hungary, Leopard fossils dating to the Late Pleistocene were found in Biśnik Cave in south-central Poland. The oldest known leopard fossils excavated in Europe are about 600,000 years old and were found in the Grotte du Vallonnet in France and near Mauer in Germany. Four European Pleistocene leopard subspecies were proposed. P. p. begoueni from the beginning of the Early Pleistocene was replaced about by P. p. sickenbergi, which in turn was replaced by P. p. antiqua around 0.3 million years ago. The most recent, P. p. spelaea, appeared at the beginning of the Late Pleistocene and survived until about 24,000 years ago in several parts of Europe. Leopard fossils dating to the Pleistocene were also excavated in the Japanese archipelago.
In 1953, a male leopard and a lioness were crossbred in Hanshin Park in Nishinomiya, Japan. Their offspring known as a leopon was born in 1959 and 1961, all cubs were spotted and bigger than a juvenile leopard. Attempts to mate a leopon with a tigress were unsuccessful.
Distribution and habitat
The leopard has the largest distribution of all wild cats, occurring widely in Africa, the Caucasus and Asia, although populations are fragmented and declining. It is considered to be extirpated in North Africa. It inhabits foremost savanna and rainforest, and areas where grasslands, woodlands, and riverine forests remain largely undisturbed. In sub-Saharan Africa, it is still numerous and surviving in marginal habitats where other large cats have disappeared. There is considerable potential for human-leopard conflict due to leopards preying on livestock.
Leopard populations on the Arabian Peninsula are small and fragmented. In southeastern Egypt, a leopard killed in 2017 was the first record in this area in 65 years. In western and central Asia, it avoids deserts, areas with long snow cover and proximity to urban centres.
In the Indian subcontinent, the leopard is still relatively abundant, with greater numbers than those of other Panthera species. As of 2020, the leopard population within forested habitats in India's tiger range landscapes was estimated at 12,172 to 13,535 individuals. Surveyed landscapes included elevations below 2,600 m (8,500 ft) in the Shivalik Hills and Gangetic plains, Central India and Eastern Ghats, Western Ghats, the Brahmaputra River basin and hills in Northeast India. Some leopard populations in the country live quite close to human settlements and even in semi-developed areas. Although adaptable to human disturbances, leopards require healthy prey populations and appropriate vegetative cover for hunting for prolonged survival and thus rarely linger in heavily developed areas. Due to the leopard's stealth, people often remain unaware that it lives in nearby areas.
In Nepal's Kanchenjunga Conservation Area, a melanistic leopard was photographed at an elevation of 4,300 m (14,100 ft) by a camera trap in May 2012. In Sri Lanka, leopards were recorded in Yala National Park and in unprotected forest patches, tea estates, grasslands, home gardens, pine and eucalyptus plantations. In Myanmar, leopards were recorded for the first time by camera traps in the hill forests of Myanmar's Karen State. The Northern Tenasserim Forest Complex in southern Myanmar is considered a leopard stronghold. In Thailand, leopards are present in the Western Forest Complex, Kaeng Krachan-Kui Buri, Khlong Saeng-Khao Sok protected area complexes and in Hala Bala Wildlife Sanctuary bordering Malaysia. In Peninsular Malaysia, leopards are present in Belum-Temengor, Taman Negara and Endau-Rompin National Parks. In Laos, leopards were recorded in Nam Et-Phou Louey National Biodiversity Conservation Area and Nam Kan National Protected Area. In Cambodia, leopards inhabit deciduous dipterocarp forest in Phnom Prich Wildlife Sanctuary and Mondulkiri Protected Forest. In southern China, leopards were recorded only in the Qinling Mountains during surveys in 11 nature reserves between 2002 and 2009.
In Java, leopards inhabit dense tropical rainforests and dry deciduous forests at elevations from sea level to 2,540 m (8,330 ft). Outside protected areas, leopards were recorded in mixed agricultural land, secondary forest and production forest between 2008 and 2014.
Behaviour and ecology
The leopard is a solitary and territorial animal. It is typically shy and alert when crossing roadways and encountering oncoming vehicles, but may be emboldened to attack people or other animals when threatened. Adults associate only in the mating season. Females continue to interact with their offspring even after weaning and have been observed sharing kills with their offspring when they can not obtain any prey. They produce a number of vocalizations, including growls, snarls, meows, and purrs. The roaring sequence in leopards consists mainly of grunts, also called "sawing", as it resembles the sound of sawing wood. Cubs call their mother with a urr-urr sound.
The whitish spots on the back of its ears are thought to play a role in communication. It has been hypothesized that the white tips of their tails may function as a 'follow-me' signal in intraspecific communication. However, no significant association were found between a conspicuous colour of tail patches and behavioural variables in carnivores.
Leopards are active mainly from dusk till dawn and rest for most of the day and for some hours at night in thickets, among rocks or over tree branches. Leopards have been observed walking 1–25 km (0.62–15.53 mi) across their range at night; they may even wander up to 75 km (47 mi) if disturbed. In some regions, they are nocturnal. In western African forests, they have been observed to be largely diurnal and hunting during twilight, when their prey animals are active; activity patterns vary between seasons.
Leopards can climb trees very skillfully, often rest on tree branches and descend from trees headfirst. They can run at over 58 km/h (36 mph), leap over 6 m (20 ft) horizontally, and jump up to 3 m (9.8 ft) vertically.
In Kruger National Park, most leopards tend to keep 1 km (0.62 mi) apart. Males interact with their partners and cubs at times, and exceptionally this can extend beyond to two generations. Aggressive encounters are rare, typically limited to defending territories from intruders. In a South African reserve, a male was wounded in a male–male territorial battle over a carcass.
Males occupy home ranges that often overlap with a few smaller female home ranges, probably as a strategy to enhance access to females. In the Ivory Coast, the home range of a female was completely enclosed within a male's. Females live with their cubs in home ranges that overlap extensively, probably due to the association between mothers and their offspring. There may be a few other fluctuating home ranges belonging to young individuals. It is not clear if male home ranges overlap as much as those of females do. Individuals try to drive away intruders of the same sex.
A study of leopards in the Namibian farmlands showed that the size of home ranges was not significantly affected by sex, rainfall patterns or season; the higher the prey availability in an area, the greater the leopard population density and the smaller the size of home ranges, but they tend to expand if there is human interference. Sizes of home ranges vary geographically and depending on habitat and availability of prey. In the Serengeti, males have home ranges of 33–38 km2 (13–15 sq mi) and females of 14–16 km2 (5.4–6.2 sq mi); but males in northeastern Namibia of 451 km2 (174 sq mi) and females of 188 km2 (73 sq mi). They are even larger in arid and montane areas. In Nepal's Bardia National Park, male home ranges of 48 km2 (19 sq mi) and female ones of 5–7 km2 (1.9–2.7 sq mi) are smaller than those generally observed in Africa.
Hunting and diet
The leopard is a carnivore that prefers medium-sized prey with a body mass ranging from 10–40 kg (22–88 lb). Prey species in this weight range tend to occur in dense habitat and to form small herds. Species that prefer open areas and have well-developed anti-predator strategies are less preferred. More than 100 prey species have been recorded. The most preferred species are ungulates, such as impala (Aepyceros melampus), bushbuck (Tragelaphus scriptus), common duiker (Sylvicapra grimmia) and chital (Axis axis). Primates preyed upon include white-eyelid mangabeys (Cercocebus sp.), guenons (Cercopithecus sp.) and gray langurs (Semnopithecus sp.). Leopards also kill smaller carnivores like black-backed jackal (Lupulella mesomelas), bat-eared fox (Otocyon megalotis), genet (Genetta sp.) and cheetah.
The largest prey killed by a leopard was reportedly a male eland weighing 900 kg (2,000 lb). A study in Wolong National Nature Reserve in southern China demonstrated variation in the leopard's diet over time; over the course of seven years, the vegetative cover receded, and leopards opportunistically shifted from primarily consuming tufted deer (Elaphodus cephalophus) to pursuing bamboo rats (Rhizomys sinense) and other smaller prey.
The leopard depends mainly on its acute senses of hearing and vision for hunting. It primarily hunts at night in most areas. In western African forests and Tsavo National Park, they have also been observed hunting by day. They usually hunt on the ground. In the Serengeti, they have been observed to ambush prey by jumping down on it from trees.
The animal stalks its prey and tries to approach as closely as possible, typically within 5 m (16 ft) of the target, and, finally, pounces on it and kills it by suffocation. It kills small prey with a bite to the back of the neck, but holds larger animals by the throat and strangles them. It caches kills up to 2 km (1.2 mi) apart. It is able to take large prey due to its powerful jaw muscles, and is therefore strong enough to drag carcasses heavier than itself up into trees; an individual was seen to haul a young giraffe weighing nearly 125 kg (276 lb) up 5.7 m (18 ft 8 in) into a tree. It eats small prey immediately, but drags larger carcasses over several hundred metres and caches it safely in trees, bushes or even caves; this behaviour allows the leopard to store its prey away from rivals, and offers it an advantage over them. The way it stores the kill depends on local topography and individual preferences, varying from trees in Kruger National Park to bushes in the plain terrain of the Kalahari.
Average daily consumption rates of 3.5 kg (7 lb 11 oz) were estimated for males and of 2.8 kg (6 lb 3 oz) for females. In the southern Kalahari Desert, leopards meet their water requirements by the bodily fluids of prey and succulent plants; they drink water every two to three days and feed infrequently on moisture-rich plants such as gemsbok cucumbers (Acanthosicyos naudinianus), watermelon (Citrullus lanatus) and Kalahari sour grass (Schmidtia kalahariensis).
Enemies and competitors
In parts of its global range, the leopard is sympatric with other large predators such as the tiger (Panthera tigris), lion (P. leo), cheetah, spotted hyena (Crocuta crocuta), striped hyena (Hyaena hyaena), brown hyena (Parahyaena brunnea), African wild dog (Lycaon pictus), dhole (Cuon alpinus), wolf (Canis lupus) and up to five bear species. Some of these species steal its kills, kill its cubs and even kill adult leopards. Leopards retreat up a tree in the face of direct aggression, and were observed when killing or preying on smaller competitors such as black-backed jackal, African civet (Civettictis civetta), caracal (Caracal caracal) and African wildcat (Felis lybica). Leopards generally seem to avoid encounters with adult bears, but kill vulnerable bear cubs. In Sri Lanka, a few recorded vicious fights between leopards and sloth bears (Melursus ursinus) apparently result in both animals winding up either dead or grievously injured.
While interspecies killing of full-grown leopards is generally rare, given the opportunity, both tiger and lion readily kill and consume both young and adult leopards. In the Kalahari Desert, leopards frequently lose kills to brown hyenas, if the leopard is unable to move the kill into a tree. Single brown hyenas have been observed charging at and displacing male leopards from kills. Lions occasionally fetch leopard kills from trees.
Resource partitioning occurs where leopards share their range with tigers. Leopards tend to take smaller prey, usually less than 75 kg (165 lb), where tigers are present. In areas where leopard and tiger are sympatric, coexistence is reportedly not the general rule, with leopards being few where tigers are numerous. Tigers appear to inhabit the deep parts of a forest while leopards are pushed closer to the fringes. In tropical forests, leopards do not always avoid the larger cats by hunting at different times. With relatively abundant prey and differences in the size of prey selected, tigers and leopards seem to successfully coexist without competitive exclusion or interspecies dominance hierarchies that may be more common to the leopard's co-existence with the lion in savanna habitats.
Nile crocodiles (Crocodylus niloticus) prey on leopards occasionally. One large adult leopard was grabbed and consumed by a large crocodile while attempting to hunt along a bank in Kruger National Park. Mugger crocodiles (Crocodylus palustris) reportedly killed an adult leopard in Rajasthan. An adult leopard was recovered from the stomach of a 5.5 m (18 ft 1 in) Burmese python (Python bivittatus). In Serengeti National Park, troops of 30–40 olive baboons (Papio anubis) were observed while mobbing and attacking a female leopard and her cubs.
Reproduction and life cycle
In some areas, leopards mate all year round. In Manchuria and Siberia, they mate during January and February. The female's estrous cycle lasts about 46 days, and she usually is in heat for 6–7 days. The generation length of the leopard is 9.3 years. Gestation lasts for 90 to 105 days. Cubs are usually born in a litter of 2–4 cubs. Mortality of cubs is estimated at 41–50% during the first year.
Females give birth in a cave, crevice among boulders, hollow tree or thicket. Cubs are born with closed eyes, which open four to nine days after birth. The fur of the young tends to be longer and thicker than that of adults. Their pelage is also more gray in colour with less defined spots. Around three months of age, the young begin to follow the mother on hunts. At one year of age, cubs can probably fend for themselves, but remain with the mother for 18–24 months.
The leopard is listed on CITES Appendix I, and trade is restricted to skins and body parts of 2,560 individuals in 11 sub-Saharan countries. The leopard is primarily threatened by habitat fragmentation and conversion of forest to agriculturally used land, which lead to a declining natural prey base, human–wildlife conflict with livestock herders and high leopard mortality rates. It is also threatened by trophy hunting and poaching.
Between 2002 and 2012, at least four leopards were estimated to have been poached per week in India for the illegal wildlife trade of its skins and bones. In spring 2013, 37 leopard skins were found during a 7-week long market survey in major Moroccan cities. In 2014, 43 leopard skins were detected during two surveys in Morocco. Vendors admitted to have imported skins from sub-Saharan Africa.
Surveys in the Central African Republic's Chinko area revealed that the leopard population decreased from 97 individuals in 2012 to 50 individuals in 2017. In this period, transhumant pastoralists from the border area with Sudan moved in the area with their livestock. Rangers confiscated large amounts of poison in the camps of livestock herders who were accompanied by armed merchants. They engaged in poaching large herbivores, sale of bushmeat and trading leopard skins in Am Dafok.
In Java, the leopard is threatened by illegal hunting and trade. Between 2011 and 2019, body parts of 51 Javan leopards were seized including six live individuals, 12 skins, 13 skulls, 20 canines and 22 claws.
Leopards have featured in art, mythology and folklore of many countries. In Greek mythology, it was a symbol of the god Dionysus, who was depicted wearing leopard skin and using leopards as means of transportation. In one myth, the god was captured by pirates but two leopards rescued him. During the Benin Empire, the leopard was commonly represented on engravings and sculptures and was used to symbolise the power of the king or oba, since the leopard was considered the king of the forest. The Ashanti also used the leopard as a symbol of leadership, and only the king was permitted to have a ceremonial leopard stool. Some African cultures considered the leopard to be a smarter, better hunter than the lion and harder to kill.
In Rudyard Kipling's "How the Leopard Got His Spots", one of his Just So Stories, a leopard with no spots in the High Veldt lives with his hunting partner, the Ethiopian. When they set off to the forest, the Ethiopian changed his brown skin, and the leopard painted spots on his skin. A leopard played an important role in the 1938 Hollywood film Bringing Up Baby. African chiefs, European queens, Hollywood actors and burlesque dancers wore coats made of leopard skins.
The leopard is a frequently used in heraldry, most commonly as passant. The heraldic leopard lacks spots and sports a mane, making it visually almost identical to the heraldic lion, and the two are often used interchangeably. Naturalistic leopard-like depictions appear on the coat of arms of Benin, Malawi, Somalia, the Democratic Republic of the Congo and Gabon, the last of which uses a black panther.
Attacks on people
The Leopard of Rudraprayag killed more than 125 people; the Panar Leopard was thought to have killed more than 400 people. Both were shot by British hunter Jim Corbett. The spotted devil of Gummalapur killed about 42 people in Karnataka, India.
The Ancient Romans kept leopards in captivity to be slaughtered in hunts as well as be used in executions of criminals. In Benin, leopards were kept and paraded as mascots, totems and sacrifices to deities. Several leopards were kept in a menagerie established by King John of England at the Tower of London in the 13th century; around 1235, three of these animals were given to Henry III by Holy Roman Emperor Frederick II. In modern times, leopards have been trained and tamed in circuses.
- Black panther – Variant of leopard and jaguar
- Leopard pattern
- List of largest cats
- Panther (legendary creature)
- Wozencraft, W. C. (2005). "Species Panthera pardus". In Wilson, D. E.; Reeder, D. M. (eds.). Mammal Species of the World: A Taxonomic and Geographic Reference (3rd ed.). Johns Hopkins University Press. p. 547. ISBN 978-0-8018-8221-0. OCLC 62265494.
- Ghezzo, E. & Rook, L. (2015). "The remarkable Panthera pardus (Felidae, Mammalia) record from Equi (Massa, Italy): taphonomy, morphology, and paleoecology". Quaternary Science Reviews. 110 (110): 131–151. doi:10.1016/j.quascirev.2014.12.020.
- Stein, A.B.; Athreya, V.; Gerngross, P.; Balme, G.; Henschel, P.; Karanth, U.; Miquelle, D.; Rostro-Garcia, S.; Kamler, J. F.; Laguardia, A.; Khorozyan, I. & Ghoddousi, A. (2020) [amended version of 2019 assessment]. "Panthera pardus". IUCN Red List of Threatened Species. 2020: e.T15954A163991139. doi:10.2305/IUCN.UK.2020-1.RLTS.T15954A163991139.en. Retrieved 15 January 2022.
- Kitchener, A. C.; Breitenmoser-Würsten, C.; Eizirik, E.; Gentry, A.; Werdelin, L.; Wilting, A.; Yamaguchi, N.; Abramov, A. V.; Christiansen, P.; Driscoll, C.; Duckworth, J. W.; Johnson, W.; Luo, S.-J.; Meijaard, E.; O’Donoghue, P.; Sanderson, J.; Seymour, K.; Bruford, M.; Groves, C.; Hoffmann, M.; Nowell, K.; Timmons, Z. & Tobe, S. (2017). "A revised taxonomy of the Felidae: The final report of the Cat Classification Task Force of the IUCN Cat Specialist Group" (PDF). Cat News (Special Issue 11): 73–75.
- Jacobson, A. P.; Gerngross, P.; Lemeris, J. R. Jr.; Schoonover, R. F.; Anco, C.; Breitenmoser-Würsten, C.; Durant, S. M.; Farhadinia, M. S.; Henschel, P.; Kamler, J. F.; Laguardia, A.; Rostro-García, S.; Stein, A. B. & Dollar, L. (2016). "Leopard (Panthera pardus) status, distribution, and the research efforts across its range". PeerJ. 4: e1974. doi:10.7717/peerj.1974. PMC 4861552. PMID 27168983.
- Williams, S. T.; Williams, K. S.; Lewis, B. P. & Hill, R. A. (2017). "Population dynamics and threats to an apex predator outside protected areas: implications for carnivore management". Royal Society Open Science. 4 (4): 161090. Bibcode:2017RSOS....461090W. doi:10.1098/rsos.161090. PMC 5414262. PMID 28484625.
- Nowell, K. & Jackson, P. (1996). "Leopard Panthera pardus (Linnaeus, 1758)". Wild Cats: status survey and conservation action plan. Gland, Switzerland: IUCN/SSC Cat Specialist Group. Archived from the original on 2014-02-22.
- Volmer, R.; Hölzchen, E.; Wurster, A.; Ferreras, M.R. & Hertler, C. (2017). "Did Panthera pardus (Linnaeus, 1758) become extinct in Sumatra because of competition for prey? Modeling interspecific competition within the Late Pleistocene carnivore guild of the Padang Highlands, Sumatra". Palaeogeography, Palaeoclimatology, Palaeoecology. 487: 175–186. Bibcode:2017PPP...487..175V. doi:10.1016/j.palaeo.2017.08.032.
- Chi T.-C.; Gan Y.; Yang T.-R. & Chang, C.-H. (2021). "First report of leopard fossils from a limestone cave in Kenting area, southern Taiwan". PeerJ. 9: e12020. doi:10.7717/peerj.12020. PMC 8388558. PMID 34513335.
- Izawa, M. & Nakanishi, N. (2015). "Felidae". In Ohdachi, S. D.; Ishibashi, Y.; Iwasa, M. A. & Saitoh, T. (eds.). The Wild Mammals of Japan (Second ed.). Kyoto: Shoukadoh Book Sellers and the Mammalogical Society of Japan. pp. 226−231. ISBN 978-4-87974-691-7.
- Lewis, C. T. & Short, C. (1879). "lěǒpardus". A Latin Dictionary. Oxford: Clarendon Press. p. 1069.
- Liddell, H. G. & Scott, R. (1889). "λέο-πάρδος". A Greek–English Lexicon. Oxford: Clarendon Press. p. 884.
- Partridge, E. (1983). Origins: A Short Etymological Dictionary of Modern English. New York: Greenwich House. p. 349. ISBN 978-0-517-41425-5.
- Nicholas, N. (1999). "A conundrum of cats: pards and their relatives in Byzantium". Greek, Roman, and Byzantine Studies. 40: 253–298. S2CID 56160515.
- Lewis, C. T. & Short, C. (1879). "panthera". A Latin Dictionary. Oxford: Clarendon Press. p. 1298.
- Lewis, C. T. & Short, C. (1879). "pardus". A Latin Dictionary. Oxford: Clarendon Press. p. 1302.
- Mills, M. G. L. (2005). "Subfamily Pantherinae". In Skinner, J. D.; Chimimba, C. T. (eds.). The mammals of the southern African subregion (Third ed.). Cambridge: Cambridge University Press. pp. 385–396. ISBN 9780521844185.
- Mivart, St. G. J. (1900). "Different kind of Cats". The Cat: An Introduction to the Study of Backboned Animals, Especially Mammals. London: John Murray. pp. 391–439.
- Pocook, R. I. (1932). "The Leopards of Africa". Proceedings of the Zoological Society of London. 102 (2): 543–591. doi:10.1111/j.1096-3642.1932.tb01085.x.
- Schütze, H. (2002). Field Guide to the Mammals of the Kruger National Park. Cape Town, South Africa: Struik Publishers. pp. 92–93. ISBN 978-1-86872-594-6.
- Menon, V. (2014). Indian Mammals: A Field Guide. Gurgaon, India: Hachette. ISBN 978-93-5009-761-8.
- Allen, W. L.; Cuthill, I. C.; Scott-Samuel, N. E. & Baddeley, R. (2010). "Why the leopard got its spots: relating pattern development to ecology in felids". Proceedings of the Royal Society B. 278 (1710): 1373–1380. doi:10.1098/rspb.2010.1734. PMC 3061134. PMID 20961899.
- Hoath, R. (2009). "Leopard Panthera pardus (Linnaeus, 1758)". Field Guide to the Mammals of Egypt. Cairo, Egypt: American University in Cairo Press. pp. 106–107. ISBN 978-977-416-254-1.
- Estes, R. (1991). "Leopard Panthera pardus". The Behavior Guide to African Mammals, Including Hoofed Mammals, Carnivores, Primates. Los Angeles: The University of California Press. pp. 366–369. ISBN 978-0-520-08085-0.
- Stein, A. B. & Hayssen, V. (2010). "Panthera pardus (Carnivora: Felidae)". Mammalian Species. 45 (900): 30–48. doi:10.1644/900.1. S2CID 44839740.
- Heptner, V. G. & Sludskii, A. A. (1992) . "Bars (leopard)". Mlekopitajuščie Sovetskogo Soiuza. Moskva: Vysšaia Škola [Mammals of the Soviet Union, Volume II, Part 2]. Washington DC: Smithsonian Institution and the National Science Foundation. pp. 203–273. ISBN 978-90-04-08876-4.
- Tanomtong, A.; Khunsook, S.; Keawmad, P. & Pintong, K. (2008). "Cytogenetic study of the leopard, Panthera pardus (Carnivora, Felidae) by conventional staining, G-banding and high-resolution staining technique". Cytologia. 73 (1): 81–90. doi:10.1508/cytologia.73.81.
- Nowak, R. M. (1999). "Panthera pardus (Leopard)". Walker's Mammals of the World (Sixth ed.). Baltimore, USA: Johns Hopkins University Press. pp. 828–831. ISBN 978-0-8018-5789-8.
- Pease, A. E. (1913). "Of dangerous game". The Book of the Lion. London: John Murray. pp. 46–68.
- Burnie, D. & Wilson, D. E., eds. (2001). Animal: The Definitive Visual Guide to the World's Wildlife. DK Adult. ISBN 978-0-7894-7764-4.
- "Is this the longest leopard in India?". The Times of India. 2016.
- "Leopard shot in Bilaspur turns out to be a record breaker". The Tribune Trust. 2016.
- Prater, S. H. (1921). "Record Panther Skull (P. p. pardus)". The Journal of the Bombay Natural History Society. XXVII (Part IV): 933–935.
- Eizirik, E.; Yuhki, N.; Johnson, W. E.; Menotti-Raymond, M.; Hannah, S. S.; O'Brien, S. J. (2003). "Molecular genetics and evolution of melanism in the cat family" (PDF). Current Biology. 13 (5): 448–453. doi:10.1016/S0960-9822(03)00128-3. PMID 12620197. S2CID 19021807. Archived from the original (PDF) on 2013-05-06.
- Robinson, R. (1970). "Inheritance of the black form of the leopard Panthera pardus". Genetica. 41 (1): 190–197. doi:10.1007/BF00958904. PMID 5480762. S2CID 5446868.
- da Silva L. G., K.; Kawanishi, K.; Henschel P.; Kittle, A.; Sanei, A.; Reebin, A.; Miquelle, D.; Stein, A. B.; Watson, A.; Kekule, L. B.; Machado, R. B. & Eizirik, E. (2017). "Mapping black panthers: Macroecological modeling of melanism in leopards (Panthera pardus)". PLOS ONE. 12 (4): e0170378. Bibcode:2017PLoSO..1270378D. doi:10.1371/journal.pone.0170378. PMC 5381760. PMID 28379961.
- Kawanishi, K.; Sunquist, M. E.; Eizirik, E.; Lynam, A. J.; Ngoprasert, D.; Wan Shahruddin, W. N.; Rayan, D. M.; Sharma, D. S. K. & Steinmetz, R. (2010). "Near fixation of melanism in leopards of the Malay Peninsula". Journal of Zoology. 282 (3): 201–206. doi:10.1111/j.1469-7998.2010.00731.x.
- Shuker, K. P. N. (2003). The Beasts that Hide from Man : Seeking the World's Last Undiscovered Animals. New York, USA: Paraview Press. p. 273. ISBN 978-1-931044-64-6.
- Divyabhanusinh (1993). "On mutant leopards Panthera pardus from India". Journal of the Bombay Natural History Society. 90 (1): 88−89.
- Pirie, T. J.; Thomas, R. L. & Fellowes, M. D. E. (2016). "Erythristic leopards Panthera pardus in South Africa". Bothalia. 46 (1): 1–5. doi:10.4102/abc.v46i1.2034.
- Linnaeus, C. (1758). "Felis pardus". Caroli Linnæi Systema naturæ per regna tria naturæ, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Vol. Tomus I (decima, reformata ed.). Holmiae: Laurentius Salvius. p. 41−42. (in Latin)
- Oken, L. (1816). "1. Art, Panthera". Lehrbuch der Zoologie. 2. Abtheilung. Jena: August Schmid & Comp. p. 1052.
- Ellerman, J. R.; Morrison-Scott, T. C. S. (1966). Checklist of Palaearctic and Indian mammals 1758 to 1946 (Second ed.). London: British Museum of Natural History. pp. 315–317.
- Allen, J. A. (1902). "Mammal names proposed by Oken in his 'Lehrbuch der Zoologie'" (PDF). Bulletin of the American Museum of Natural History. 16 (27): 373−379.
- Pocock, R. I. (1917). "The Classification of existing Felidae". The Annals and Magazine of Natural History. Series 8. XX: 329–350. doi:10.1080/00222931709487018.
- Pocock, R. I. (1939). "Panthera pardus". The Fauna of British India, including Ceylon and Burma. Mammalia: Volume 1. London: Taylor and Francis. pp. 222–239.
- Miththapala, S.; Seidensticker, J. & O'Brien, S. J. (1996). "Phylogeographic subspecies recognition in leopards (Panthera pardus): molecular genetic variation" (PDF). Conservation Biology. 10 (4): 1115–1132. doi:10.1046/j.1523-1739.1996.10041115.x.
- Uphyrkina, O.; Johnson, E. W.; Quigley, H.; Miquelle, D.; Marker, L.; Bush, M. & O'Brien, S. J. (2001). "Phylogenetics, genome diversity and origin of modern leopard, Panthera pardus" (PDF). Molecular Ecology. 10 (11): 2617–2633. doi:10.1046/j.0962-1083.2001.01350.x. PMID 11883877. S2CID 304770. Archived (PDF) from the original on 2011-09-10.
- Meyer, F. A. A. (1794). "Über de la Metheries schwarzen Panther". Zoologische Annalen. Erster Band. Weimar: Im Verlage des Industrie-Comptoirs. pp. 394–396.
- Laguardia, A.; Kamler, J. F.; Li, S.; Zhang, C.; Zhou, Z.; Shi, K. (2017). "The current distribution and status of leopards Panthera pardus in China". Oryx. 51 (1): 153−159. doi:10.1017/S0030605315000988.
- Cuvier, G. (1809). "Recherches sur les espėces vivantes de grands chats, pour servir de preuves et d'éclaircissement au chapitre sur les carnassiers fossils". Annales du Muséum National d'Histoire Naturelle. Tome XIV: 136–164.
- Hemprich, W.; Ehrenberg, C. G. (1830). "Felis, pardus?, nimr". In Dr. C. G. Ehrenberg (ed.). Symbolae Physicae, seu Icones et Descriptiones Mammalium quae ex Itinere per Africam Borealem et Asiam Occidentalem Friderici Guilelmi Hemprich et Christiani Godofredi Ehrenberg. Decas Secunda. Zoologica I. Mammalia II. Berolini: Officina Academica. pp. Plate 17.
- Spalton, J. A. & Al Hikmani, H. M. (2006). "The Leopard in the Arabian Peninsula – Distribution and Subspecies Status" (PDF). Cat News (Special Issue 1): 4–8. Archived (PDF) from the original on 2015-06-19.
- Valenciennes, A. (1856). "Sur une nouvelles espèce de Panthère tué par M. Tchihatcheff à Ninfi, village situé à huit lieues est de Smyrne". Comptes Rendus Hebdomadaires des Séances de l'Académie des Sciences. 42: 1035–1039.
- Khorozyan, I. G.; Gennady, F.; Baryshnikov, G. F. & Abramov, A. V. (2006). "Taxonomic status of the leopard, Panthera pardus (Carnivora, Felidae) in the Caucasus and adjacent areas". Russian Journal of Theriology. 5 (1): 41–52. doi:10.15298/rusjtheriol.05.1.06.
- Schlegel, H. (1857). "Felis orientalis". Handleiding Tot de Beoefening der Dierkunde, Ie Deel. Breda: Boekdrukkerij van Nys. p. 23.
- Gray, J. E. (1862). "Description of some new species of Mammalia". Proceedings of the Royal Zoological Society of London. 30: 261−263, plate XXXIII. doi:10.1111/j.1469-7998.1862.tb06524.x.
- Pocock, R. I. (1930). "The Panthers and Ounces of Asia". Journal of the Bombay Natural History Society. 34 (2): 307–336.
- Deraniyagala, P. E. P. (1956). "The Ceylon leopard, a distinct subspecies". Spolia Zeylanica. 28: 115–116.
- Anco, C.; Kolokotronis, S. O.; Henschel, P.; Cunningham, S. W.; Amato, G. & Hekkala, E. (2017). "Historical mitochondrial diversity in African leopards (Panthera pardus) revealed by archival museum specimens". Mitochondrial DNA Part A. 29 (3): 455–473. doi:10.1080/24701394.2017.1307973. PMID 28423965. S2CID 4348541.
- Johnson, W. E.; Eizirik, E.; Pecon-Slattery, J.; Murphy, W. J.; Antunes, A.; Teeling, E. & O'Brien, S. J. (2006). "The late Miocene radiation of modern Felidae: a genetic assessment". Science. 311 (5757): 73–77. Bibcode:2006Sci...311...73J. doi:10.1126/science.1122277. PMID 16400146. S2CID 41672825.
- Werdelin, L.; Yamaguchi, N.; Johnson, W. E. & O'Brien, S. J. (2010). "Phylogeny and evolution of cats (Felidae)". In Macdonald, D. W. & Loveridge, A. J. (eds.). Biology and Conservation of Wild Felids. Oxford, UK: Oxford University Press. pp. 59–82. ISBN 978-0-19-923445-5.
- Davis, B. W.; Li, G. & Murphy, W. J. (2010). "Supermatrix and species tree methods resolve phylogenetic relationships within the big cats, Panthera (Carnivora: Felidae)" (PDF). Molecular Phylogenetics and Evolution. 56 (1): 64–76. doi:10.1016/j.ympev.2010.01.036. PMID 20138224. Archived from the original (PDF) on 2016-03-05.
- Mazák, J. H.; Christiansen, P.; Kitchener, A. C. & Goswami, A. (2011). "Oldest known pantherine skull and evolution of the tiger". PLOS ONE. 6 (10): e25483. Bibcode:2011PLoSO...625483M. doi:10.1371/journal.pone.0025483. PMC 3189913. PMID 22016768.
- Bininda-Emonds, O. R. P.; Decker-Flum, D. M. & Gittleman, J. L. (2001). "The utility of chemical signals as phylogenetic characters: an example from the Felidae". Biological Journal of the Linnean Society. 72 (1): 1–15. doi:10.1111/j.1095-8312.2001.tb01297.x.
- Tseng, Z. J.; Wang, X.; Slater, G. J.; Takeuchi, G. T.; Li, Q.; Liu, J. & Xie, G. (2014). "Himalayan fossils of the oldest known pantherine establish ancient origin of big cats". Proceedings of the Royal Society B: Biological Sciences. 281 (1774): 20132686. doi:10.1098/rspb.2013.2686. PMC 3843846. PMID 24225466.
- Li, G.; Davis, B. W.; Eizirik, E. & Murphy, W. J. (2016). "Phylogenomic evidence for ancient hybridization in the genomes of living cats (Felidae)". Genome Research. 26 (1): 1–11. doi:10.1101/gr.186668.114. PMC 4691742. PMID 26518481.
- Wilting, A.; Patel, R.; Pfestorf, H.; Kern, C.; Sultan, K.; Ario, A.; Peñaloza, F.; Kramer‐Schadt, S.; Radchuk, V.; Foerster, D.W. & Fickel, J. (2016). "Evolutionary history and conservation significance of the Javan leopard Panthera pardus melas". Journal of Zoology. 299 (4): 239–250. doi:10.1111/jzo.12348.
- Schmid, E. (1940). "Variationstatistische Untersuchungen am Gebiss pleistozäner und rezenter Leoparden und anderer Feliden". Zeitschrift für Säugetierkunde. 15: 1–179.
- Marciszak, A. & Stefaniak, K. (2010). "Two forms of cave lion: Middle Pleistocene Panthera spelaea fossilis Reichenau, 1906 and Upper Pleistocene Panthera spelaea spelaea Goldfuss, 1810 from the Bísnik Cave, Poland". Neues Jahrbuch für Geologie und Paläontologie - Abhandlungen. 258 (3): 339–351. doi:10.1127/0077-7749/2010/0117.
- Diedrich, C. G. (2013). "Late Pleistocene leopards across Europe – northernmost European German population, highest elevated records in the Swiss Alps, complete skeletons in the Bosnia Herzegowina Dinarids and comparison to the Ice Age cave art". Quaternary Science Reviews. 76: 167–193. Bibcode:2013QSRv...76..167D. doi:10.1016/j.quascirev.2013.05.009.
- Kawata, K. (2001). "Zoological gardens of Japan". In Kisling, V.N. (ed.). Zoo and Aquarium History : Ancient Animal Collections to Zoological Gardens. Boca Raton, Florida: CRC Press. pp. 295–329. ISBN 978-0-8493-2100-9.
- Murphey, R. (1951). "The Decline of North Africa Since the Roman Occupation: Climatic or Human?" (PDF). Annals of the Association of American Geographers. XLI (2): 116–132. doi:10.1080/00045605109352048. Archived (PDF) from the original on 2006-09-14.
- Pirie, T. J.; Thomas, R. L. & Fellowes, M. D. E. (2017). "Increasing game prices may alter farmers' behaviours towards leopards (Panthera pardus) and other carnivores in South Africa". PeerJ. 5: e3369. doi:10.7717/peerj.3369. PMC 5452990. PMID 28584709.
- Spalton, J. A. & Al Hikmani, H. M. (2006). "The Leopard in the Arabian Peninsula – Distribution and Subspecies Status" (PDF). Cat News (Special Issue 1): 4–8. Archived (PDF) from the original on 2011-05-23.
- Judas, J.; Paillat, P.; Khoja, A. & Boug, A. (2006). "Status of the Arabian leopard in Saudi Arabia" (PDF). Cat News (Special Issue 1): 11–19. Archived (PDF) from the original on 2015-09-19.
- Al Jumaily, M.; Mallon, D. P.; Nasher, A. K. & Thowabeh, N. (2006). "Status Report on Arabian Leopard in Yemen". Cat News (Special Issue 1): 20–25.
- Soultan, A.; Attum, O.; Hamada, A.; Hatab, E. B.; Ahmed, S. E.; Eisa, A.; Al Sharif, I.; Nagy, A. & Shohdi, W. (2017). "Recent observation for leopard Panthera pardus in Egypt". Mammalia. 81 (1): 115–117. doi:10.1515/mammalia-2015-0089. S2CID 90676105.
- Gavashelishvili, A. & Lukarevskiy, V. (2008). "Modelling the habitat requirements of leopard Panthera pardus in west and central Asia". Journal of Applied Ecology. 45 (2): 579–588. doi:10.1111/j.1365-2664.2007.01432.x.
- Jhala, Y.V.; Qureshi, Q. & Yadav, S.P. (2020). Status of leopards in India, 2018. Technical Report TR/2020/16 (Report). New Delhi and Dehradun: National Tiger Conservation Authority, Government of India and Wildlife Institute of India.
- Arthreya, V. (2012). "Living with Leopards Outside Protected Areas in India". Conservation India.
- Thapa, K.; Pradhan, N. M. B.; Berker, J.; Dhakal, M.; Bhandari, A. R.; Gurung, G. S.; Rai, D. P.; Thapa, G. J.; Shrestha, S. & Singh, G. R. (2013). "High elevation record of a leopard cat in the Kangchenjunga Conservation Area, Nepal". Cat News (58): 26–27.
- Kittle, A. M.; Watson, A. C.; Chanaka Kumara, P. H. & Nimalka Sanjeewani, H. K. (2014). "Status and distribution of the leopard in the central hills of Sri Lanka". Cat News (56): 28−31.
- Kittle, A. M.; Watson, A. C.; Kumara, P. H. S. C.; Sandanayake, S. D. K. C.; Sanjeewani, H. K. N. & Fernando, T. S. P. (2014). "Notes on the diet and habitat selection of the Sri Lankan Leopard Panthera pardus kotiya (Mammalia: Felidae) in the central highlands of Sri Lanka". Journal of Threatened Taxa. 6 (9): 6214–6221. doi:10.11609/JoTT.o3731.6214-21.
- Saw Sha Bwe Moo; Froese, G.Z.L. & Gray, T.N.E. (2017). "First structured camera-trap surveys in Karen State, Myanmar, reveal high diversity of globally threatened mammals". Oryx. 52 (3): 537−543. doi:10.1017/S0030605316001113.
- Rostro-García, S.; Kamler, J. F.; Ash, E.; Clements, G. R.; Gibson, L.; Lynam, A. J.; McEwin, R.; Naing, H. & Paglia, S. (2016). "Endangered leopards: Range collapse of the Indochinese leopard (Panthera pardus delacouri) in Southeast Asia". Biological Conservation. 201: 293–300. doi:10.1016/j.biocon.2016.07.001. hdl:10722/232870.
- Johnson, A.; Vongkhamheng, C.; Hedemark, M. & Saithongdam, T. (2006). "Effects of human–carnivore conflict on tiger (Panthera tigris) and prey populations in Lao PDR" (PDF). Animal Conservation. 9 (4): 421–430. doi:10.1111/j.1469-1795.2006.00049.x. S2CID 73637721. Archived (PDF) from the original on 2017-08-10.
- Robichaud, W.; Insua-Cao; Sisomphane, P. C. & Chounnavanh, S. (2010). "Appendix 4". A scoping mission to Nam Kan National Protected Area, Lao PDR. Fauna & Flora International. pp. 33−42.
- Gray, T. N. & Phan, C. (2011). "Habitat preferences and activity patterns of the larger mammal community in Phnom Prich Wildlife Sanctuary, Cambodia". The Raffles Bulletin of Zoology. 59 (2): 311−318.
- Gray, T. N. E. (2013). "Activity patterns and home ranges of Indochinese leopard Panthera pardus delacouri in the Eastern Plains Landscape, Cambodia" (PDF). Natural History Bulletin of the Siam Society. 59: 39−47. Archived (PDF) from the original on 2016-02-22.
- Li, S.; Wang, D.; Lu, Z. & Mc Shea, W. J. (2010). "Cats living with pandas: The status of wild felids within giant panda range, China". Cat News. 52: 20–23.
- Wibisono, H. T.; Wahyudi, H. A.; Wilianto, E.; Pinondang, I. M. R.; Primajati, M.; Liswanto, D. & Linkie, M. (2018). "Identifying priority conservation landscapes and actions for the Critically Endangered Javan leopard in Indonesia: Conserving the last large carnivore in Java Island". PLOS ONE. 13 (6): e0198369. Bibcode:2018PLoSO..1398369W. doi:10.1371/journal.pone.0198369. PMC 6021038. PMID 29949588.
- Sunquist, M. E. & Sunquist, F. (2002). "Leopard Panthera pardus". Wild Cats of the World. Chicago: University of Chicago Press. pp. 318–342. ISBN 978-0-226-77999-7.
- Leyhausen, P. (1979). Cat behavior: the predatory and social behavior of domestic and wild cats. Berlin: Garland Publishing, Incorporated. p. 281. ISBN 9780824070175.
- Ortolani, A. (1999). "Spots, stripes, tail tips and dark eyes: predicting the function of carnivore colour patterns using the comparative method". Biological Journal of the Linnean Society. 67 (4): 433–476. doi:10.1111/j.1095-8312.1999.tb01942.x.
- Caro, T. (2005). "The adaptive significance of coloration in mammals". BioScience. 55 (2): 125–136. doi:10.1641/0006-3568(2005)055[0125:TASOCI]2.0.CO;2.
- Hunter, L.; Balme, G.; Walker, C.; Pretorius, K. & Rosenberg, K. (2003). "The landscape ecology of leopards (Panthera pardus) in northern KwaZulu-Natal, South Africa: a preliminary project report" (PDF). Ecological Journal. 5: 24–30. Archived from the original (PDF) on March 4, 2009.
- Spalton, J.A.; Al Hikmani, H. M.; Willis, D. & Said, A. S. B. (2006). "Critically endangered Arabian leopards Panthera pardus nimr persist in the Jabal Samhan Nature Reserve, Oman". Oryx. 40 (3): 287–294. doi:10.1017/S0030605306000743.
- Jenny, D. & Zuberbuhler, K. (2005). "Hunting behaviour in west African forest leopards". African Journal of Ecology. 43 (3): 197–200. doi:10.1111/j.1365-2028.2005.00565.x.
- Bailey, T. N. (1993). The African leopard: a study of the ecology and behaviour of a solitary felid. New York: Columbia University Press. ISBN 978-1-932846-11-9.
- Hunter, L.; Henschel, P. & Ray, J. C. (2013). "Panthera pardus Leopard". In Kingdon, J.; Happold, D.; Butynski, T.; Hoffmann, M.; Happold, M. & Kalina, J. (eds.). Mammals of Africa. London: Bloomsbury Publishing. pp. 159–168. ISBN 978-1-4081-8996-2.
- Pirie, T. J.; Thomas, R. L.; Reilly, B. K. & Fellowes, M. D. E. (2014). "Social interactions between a male leopard (Panthera pardus) and two generations of his offspring". African Journal of Ecology. 52 (4): 574–576. doi:10.1111/aje.12154.
- Jenny, D. (1996). "Spatial organization of leopards Panthera pardus in Tai National Park, Ivory Coast: Is rainforest habitat a "tropical haven"?". Journal of Zoology. 240 (3): 427–440. doi:10.1111/j.1469-7998.1996.tb05296.x.
- Marker, L. L. & Dickman, A. J. (2005). "Factors affecting leopard (Panthera pardus) spatial ecology, with particular reference to Namibian farmlands" (PDF). South African Journal of Wildlife Research. 35 (2): 105–115.
- Bertram, B. C. R. (1982). "Leopard ecology as studied by radio tracking". Symposia of the Zoological Society of London. 49: 341–352.
- Mizutani, F. & Jewell, P. A. (1998). "Home-range and movements of leopards (Panthera pardus) on a livestock ranch in Kenya". Journal of Zoology. 244 (2): 269–286. doi:10.1017/S0952836998002118.
- Stander, P. E.; Haden, P. J.; Kaqece, II. & Ghau, II. (1997). "The ecology of asociality in Namibian leopards". Journal of Zoology. 242 (2): 343–364. doi:10.1111/j.1469-7998.1997.tb05806.x.
- Odden, M. & Wegge, P. (2005). "Spacing and activity patterns of leopards Panthera pardus in the Royal Bardia National Park, Nepal". Wildlife Biology. 11 (2): 145–152. doi:10.2981/0909-6396(2005)11[145:SAAPOL]2.0.CO;2. S2CID 86140708.
- Hayward, M.W.; Henschel, P.; O'Brien, J.; Hofmeyr, M.; Balme, G. & Kerley, G. I. H. (2006). "Prey preferences of the leopard (Panthera pardus)" (PDF). Journal of Zoology. 270 (4): 298–313. doi:10.1111/j.1469-7998.2006.00139.x. Archived (PDF) from the original on 2012-11-05.
- Johnson, K. G.; Wei, W.; Reid, D. G.; Jinchu, H. (1993). "Food habits of Asiatic leopards (Panthera pardus fusca) in Wolong Reserve, Sichuan, China". Journal of Mammalogy. 74 (3): 646–650. doi:10.2307/1382285. JSTOR 1382285.
- Mills, M. G. L. & Hes, L. (1997). The Complete Book of Southern African Mammals. Cape Town, South Africa: Struik Publishers. pp. 178–180. ISBN 978-0-947430-55-9.
- Hamilton, P. H. (1976). The movements of leopards in Tsavo National Park, Kenya, as determined by radio-tracking (M.Sc. thesis). Nairobi: University of Nairobi.
- Kruuk, H. & Turner, M. (1967). "Comparative notes on predation by lion, leopard, cheetah and wild dog in the Serengeti area, East Africa". Mammalia. 31 (1): 1–27. doi:10.1515/mamm.19184.108.40.206. S2CID 84619500.
- Schaller, G. (1972). Serengeti: a kingdom of predators. New York: Knopf. ISBN 978-0-394-47242-3.
- Bothma, J. du P. (2005). "Water-use by southern Kalahari leopards" (PDF). South African Journal of Wildlife Research. 35: 131–137.
- Palomares, F. & Caro, T. M. (1999). "Interspecific killing among mammalian carnivores" (PDF). The American Naturalist. 153 (5): 492–508. doi:10.1086/303189. hdl:10261/51387. PMID 29578790. S2CID 4343007. Archived from the original (PDF) on 2019-09-29.
- Kurt, F. & Jayasuriya, A. (1968). "Notes on a dead bear". Loris (11): 182–183.
- Baskaran, N.; Sivaganesan, N. & Krishnamoorthy, J. (1997). "Food habits of sloth bear in Mudumalai Wildlife Sanctuary, Tamil Nadu, southern India". Journal of the Bombay Natural History Society. 94: 1–9.
- Seidensticker, J. (1976). "On the ecological separation between tigers and leopards" (PDF). Biotropica. 8 (4): 225–234. doi:10.2307/2989714. JSTOR 2989714.
- Johnsingh, A. J. T. (1992). "Prey selection in three large sympatric carnivores in Bandipur". Mammalia. 56 (4): 517–526. doi:10.1515/mamm.19220.127.116.117. S2CID 84997827.
- Owens, D. & Owens, M. (1980). "Hyenas of the Kalahari". Natural History. 89 (2): 50.
- Owens, M. & Owens, D. (1984). Cry of the Kalahari. Boston: Houghton Mifflin. ISBN 978-0-395-32214-7.
- Thinley, P.; Rajaratnam, R.; Lassoie, J. P.; Morreale, S. J.; Curtis, P. D.; Vernes, K.; Leki Leki; Phuntsho, S.; Dorji, T. & Dorji, P. (2018). "The ecological benefit of tigers (Panthera tigris) to farmers in reducing crop and livestock losses in the eastern Himalayas: Implications for conservation of large apex predators". Biological Conservation. 219: 119–125. doi:10.1016/j.biocon.2018.08.007.
- Karanth, U. K. & Sunquist, M. E. (2000). "Behavioural correlates of predation by tiger (Panthera tigris), leopard (Panthera pardus) and dhole (Cuon alpinus) in Nagarahole, India". Journal of Zoology. 250 (2): 255–265. doi:10.1111/j.1469-7998.2000.tb01076.x.
- Bhatnagar, C.; Mahur, M. (2010). "Observations on feeding behavior of a wild population of marsh crocodile in Baghdarrah Lake, Udaipur, Rajasthan". Reptile Rap. 10: 16–18.
- Gower, D.; Garrett, K. & Stafford, P. (2012). Snakes. Firefly Books. p. 60. ISBN 978-1-55407-802-8.
- Kiffner, C.; Ndibalema, V. & Kioko, J. (2012). "Leopard (Panthera pardus) aggregation and interactions with Olive baboons (Papio anubis) in Serengeti National Park, Tanzania". African Journal of Ecology. 51 (1): 168–171. doi:10.1111/aje.12002.
- Sadleir, R. (1966). "Notes on the Reproduction of the larger Felidae". International Zoo Yearbook. 6: 184–187. doi:10.1111/j.1748-1090.1966.tb01746.x.
- Pacifici, M.; Santini, L.; Di Marco, M.; Baisero, D.; Francucci, L.; Grottolo Marasini, G.; Visconti, P. & Rondinini, C. (2013). "Generation length for mammals". Nature Conservation (5): 87–94.
- Hemmer, H. (1976). "Gestation period and postnatal development in felids". In Eaton, R.L. (ed.). The world's cats. Vol. 3. Carnivore Research Institute, Univ. Washington, Seattle. pp. 143–165.
- Eaton, R.L. (1977). "Reproductive biology of the leopard". Zoologischer Garten. 47 (5): 329–351.
- "Leopard (Panthera pardus); Physical characteristics and distribution". Comparative Mammalian Brain Collections.
- Salisbury, S. (2014). "Roxanne, oldest spotted leopard in captivity, dies at Acreage preserve". The Palm Beach Post. Archived from the original on 2014-08-11.
- Raza, R.H.; Chauhan, D.S.; Pasha, M.K.S. & Sinha, S. (2012). Illuminating the blind spot: A study on illegal trade in Leopard parts in India (2001–2010) (PDF) (Report). New Delhi: TRAFFIC India, WWF India. Archived (PDF) from the original on 2020-09-24.
- Bergin, D. & Nijman, V. (2014). "Open, Unregulated Trade in Wildlife in Morocco's Markets". TRAFFIC Bulletin. 26 (1): 65–70.
- Bergin, D. & Nijman, V. (2015). "Potential benefits of impending Moroccan wildlife trade laws, a case study in carnivore skins". Biodiversity and Conservation. 25 (1): 199–201. doi:10.1007/s10531-015-1042-1. S2CID 34533018.
- Äbischer, T.; Ibrahim, T.; Hickisch, R.; Furrer, R. D.; Leuenberger, C. & Wegmann, D. (2020). "Apex predators decline after an influx of pastoralists in former Central African Republic hunting zones" (PDF). Biological Conservation. 241: 108326. doi:10.1016/j.biocon.2019.108326. S2CID 213766740. Archived (PDF) from the original on 2020-10-03.
- Gomez, L. & Shepherd, C.R. (2021). "The illegal exploitation of the Javan Leopard (Panthera pardus melas) and Sunda Clouded Leopard (Neofelis diardi) in Indonesia". Nature Conservation. 43 (43): 25–39. doi:10.3897/natureconservation.43.59399. S2CID 233286106.
- Morris, D. (2014). Leopard. Reaktion Books. pp. 23–24, 31–33, 62, 99, 102, 111. ISBN 9781780233185.
- "Benin: an African kingdom" (PDF). London: British Museum. Archived (PDF) from the original on 2008-08-05. Retrieved 2016-03-29.
- Kipling, R. (1902). "How the Leopard Got His Spots". Just So Stories. Macmillan.
- Haist, M. (1999). "The Lion, bloodline, and kingship". In Hassig, D. (ed.). The Mark of the Beast: The Medieval Bestiary in Art, Life, and Literature. London: Taylor & Francis. pp. 3–16. ISBN 978-0-8153-2952-7.
- Pedersen, C. F. (1971). The International Flag Book in Color. Morrow.
- Corbett, J. (1955). The Temple Tiger, and More Man-eaters of Kumaon. Oxford: Oxford University Press.
- Anderson, K. (1954). "The Spotted Devil of Gummalapur". Nine Man-Eaters and one Rogue. London: George Allen & Unwin. pp. 36–51.
- Owen, J. (2005). "Medieval Lion Skulls Reveal Secrets of Tower of London 'Zoo'". National Geographic Magazine. Retrieved 2007-09-05.
- Allsen, Thomas T. (2007). "Natural History and Cultural History: The Circulation of Hunting Leopards in Eurasia, Seventh-Seventeenth Centuries". In Mair, Victor H. (ed.). Contact and Exchange in the Ancient World. Honolulu: University of Hawai'i Press. ISBN 978-0-8248-2884-4.
- DeRuiter, D. J.; Berger, L. R. (2000). "Leopards as Taphonomic Agents in dolomitic Caves—Implications for bone Accumulations in the Hominid-bearing Deposits of South Africa". Journal of Archaeological Science. 27 (8): 665–684. doi:10.1006/jasc.1999.0470.
- Schaller, G. B. (1972). The Serengeti Lion. Chicago: University of Chicago Press. ISBN 978-0-226-73639-6.
- Sanei, A. (2007). Analysis of leopard (Panthera pardus) status in Iran (in Persian). Tehran: Sepehr Publication Center. ISBN 978-964-6123-74-8.
- Sanei, A.; Zakaria, M.; Yusof, E.; Roslan, M. (2011). "Estimation of leopard population size in a secondary forest within Malaysia's capital agglomeration using unsupervised classification of pugmarks" (PDF). Tropical Ecology. 52 (1): 209–217. Archived (PDF) from the original on 2011-10-02.
- Taylor, P.; Barrientos, S.; Dolan, C. (2005). Beyond Conservation: A Wildland Strategy. Earthscan. ISBN 978-1-84407-197-5.
- Zakaria, M.; Sanei, A. (2011). "Conservation and management prospects of the Persian and Malayan leopards". Asia Life Sciences. Supplement 7: 1–5.